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Contents lists available at ScienceDirect

Scientia Horticulturae

journal homepage: www.elsevier.com/locate/scihorti

Review

Fungus resistant varieties as a suitable alternative for organic

production: Benefits, limits, and challenges

a,∗ b,∗

Karine Pedneault , Caroline Provost

a

Centre de développement bioalimentaire du Québec, 1642 rue de la Ferme, La Pocatiere, Quebec, Canada

b

Centre de recherche agroalimentaire de Mirabel, 9850 Belle-Rivière, Mirabel, Quebec, Canada

a r t i c l e i n f o a b s t r a c t

Article history: Areas dedicated to organic wine production have significantly increased over the last few years. The

Received 5 November 2015

vast majority of organic wine is made from vinifera varieties that are highly susceptible to fungal

Received in revised form 8 March 2016

diseases and pests, making organic management difficult for growers. Depending on the growing area,

Accepted 14 March 2016

20–70% of organic growers declare issues with fungal diseases in Europe. Recently, fungus-resistant grape

Available online xxx

(FRG) varieties have been recommended as the most suitable choice in organic , especially in

areas where disease pressure necessitates high rates of fungicides. FRG varieties could contribute to

Keywords:

improved disease management in organic as well as conventional viticulture, reduce production costs

Interspecific hybrid

and decrease copper accumulation in soils. Recently, many FRG varieties presenting advantageous agro-

PIWI grape

nomic attributes and enological characteristics have been developed in North America and Europe for

Organic viticulture

Breeding conventional and sustainable farming. In this review, we present an overview of the benefits and limits

Phenolic compounds associated with FRG varieties in addition to the current knowledge regarding berry and wine composition,

Volatile compounds canopy management, and winemaking challenges and practices.

Resistance

© 2016 Elsevier B.V. All rights reserved.

Powdery mildew

Downy mildew Botrytis

Contents

1. Introduction ...... 00

2. Benefits and limits of FRG in organic viticulture ...... 00

2.1. Breeding for disease resistance and quality ...... 00

2.2. Economical and agricultural benefits of disease resistance ...... 00

2.3. Yield ...... 00

2.4. Marketing and wine quality...... 00

3. Growing FRG varieties for organic wine production ...... 00

3.1. Berry and juice composition ...... 00

3.1.1. Juice ...... 00

3.1.2. Berries ...... 00

3.2. Impacts of canopy management ...... 00

3.2.1. Managing yield and berry quality ...... 00

3.2.2. Impact on wine ...... 00

3.3. Challenges in FRG wine production ...... 00

3.3.1. Juice extraction and methanol ...... 00

Abbreviations: FRG, fungus resistant grape; M3GE, malvidin-3-glucoside equivalents; MDGE, malvidin-3,5-diglucoside equivalents; DW, dry weight; FW, fresh weight;

LC–MS/MS, liquid chromatography coupled to mass spectrometry; UPLC–MS, ultraperformance liquid chromatography coupled to mass spectrometry; HPLC-DAD, high

performance liquid chromatography coupled to diode-array detector; CE, catechin equivalent; B2E, procyanidin B2 equivalent; RE, resveratrol equivalent; TA, titratable

acidity; TSS, total soluble solids; YAN, yeast assimilable nitrogen. ∗

Corresponding authors.

E-mail addresses: [email protected], [email protected], [email protected] (K. Pedneault), [email protected] (C. Provost).

http://dx.doi.org/10.1016/j.scienta.2016.03.016

0304-4238/© 2016 Elsevier B.V. All rights reserved.

Please cite this article in press as: Pedneault, K., Provost, C., Fungus resistant grape varieties as a suitable alternative for organic wine

production: Benefits, limits, and challenges. Sci. Hortic. (2016), http://dx.doi.org/10.1016/j.scienta.2016.03.016

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3.3.2. Biogenic amines ...... 00

3.3.3. Tannins and color...... 00

3.3.4. Aroma ...... 00

3.4. Improving FRG wine quality ...... 00

4. Conclusion ...... 00

Acknowledgements...... 00

References ...... 00

1. Introduction carrying both disease-resistance genes and a significant percentage

(more than 85%) of V. vinifera in their pedigree; those are gen-

Organic wine production considerably increased over the last erally referred to as “PIWI” (from German: Pilzwiderstandsfähige,

ten years and accounts for nearly 5% of total production of today’s “disease resistant”) and are accepted as V. vinifera varieties in

wine market (Bonn et al., 2015; Willer and Lernoud, 2015). Rea- European catalogues (Sivcevˇ et al., 2010). In some cases though,

sons driving consumers toward organic wine varies from health “PIWI” indistinctly refers to both interspecific hybrids and newer

concerns to environmental awareness and interest for terroir, and “disease-resistant V. vinifera” varieties (Collective, 2008; Siegfried

the overall perception that organic are of higher value than and Temperli, 2008).

conventional ones (Bonn et al., 2015). Optimal variety selection is a key factor for successful imple-

Organic wines are expected to be free of synthetic pesticides, fer- mentation of organic grape production (Fragoulis et al., 2009;

tilizers or other synthetic inputs that could pose a higher risk than Sivcevˇ et al., 2010). Recently, FRG varieties have been rec-

conventional farming to human health or the environment (Mann ommended as the most suitable choice for organic viticulture

et al., 2010; Bonn et al., 2015). However, the vast majority of organic (Pavlousek,ˇ 2010; Sivcevˇ et al., 2010; Becker, 2013). Resistance

wine is made from varieties highly susceptible to fun- to major diseases such as powdery mildew significantly reduces

gal diseases and pests, making organic management difficult for the need for pesticides and thus represents a major advantage

growers (Wiedemann-Merdinoglu and Hoffmann, 2010). Depend- in organic farming, especially in humid areas such as Bordeaux

ing on the country origin, 20–70% of organic growers declare issues (Galbrun, 2008; Sivcevˇ et al., 2010; Wiedemann-Merdinoglu and

with botrytis and powdery mildew in Europe (Collective, 2008). Hoffmann, 2010; Fuller et al., 2014; Weigle and Carroll, 2015).

In contrast with conventional viticulture that integrates a Superficies devoted to organic wine production from FRG vari-

wide range of synthetic pesticides in pest management programs, eties are not well documented. In Germany, FRG occupied 7.9% of

organic viticulture mostly relies on sulfur- and copper-based fungi- organic vineyard surface areas in 2003 but projections were that

cides such as the Bordeaux mixture to control major diseases like 40% of the new plantings planned from 2010 to 2015 would be FRG

downy and powdery mildews, as well as a wide range of other cultivars (Sloan et al., 2010). In contrast, both old and recent FRG

diseases and insect pests (Provenzano et al., 2010). Copper-based varieties have spread extensively over the last 30 years in areas pre-

formulations have been used for more than a hundred years in senting challenging growing conditions such as wet summers and

European vineyards (Flores-Vélez et al., 1996), but copper has a cold winters with most of these varieties grown using conventional

low mobility in soil and was later found to accumulate to lev- management practices (Table 1). Very little research has been done

els that could threaten the environment (Komárek et al., 2010). to compare vineyard practices and pesticide use for FRG varieties

Indeed, copper concentration is typically much higher in vine- grown under conventional vs. organic management. In addition,

yard soils (20–665 mg/kg) compared to arable land (5–30 mg/kg) the large genetic pool found in FRG varieties makes them highly

(Besnard et al., 2001; Komárek et al., 2010). Such accumulation is variable in terms of viticulture (e.g., vigor, canopy management,

likely to occur in perennial crops like grapevines because copper berry ripening), berry composition (e.g., sugars, acids, phenolic

fungicides are continuously sprayed on the same land (Komárek compounds) and winemaking (e.g., aroma). In order to obtain a

et al., 2010). Correspondingly, vineyards located in wet areas show comprehensive understanding of FRG varieties with regards to

higher copper concentration than those from dry areas, which organic grape production, this review will present the benefits and

suffer less pressure from diseases (Komárek et al., 2010). Cop- limits of FRG for organic wine production, present their agronomic

per concentration reached 93 mg/kg in the 0–20 cm soil layer of a and oenological characteristics, and discuss challenges related to

Mediterranean organic vineyard with a dry climate, whereas values their use in wine production.

between 200 and 297 mg/kg have been reported in conventional

vineyards located in wet areas from Northern Italy (Provenzano

2. Benefits and limits of FRG in organic viticulture

et al., 2010).

The European Union recently established premium goals to

2.1. Breeding for disease resistance and quality

reduce pesticides, particularly copper, in viticulture (Rousseau

et al., 2013). One of the strategies is to shift from a treatment-

In phytopathology, “resistance” refers to the capacity of the

oriented approach to a disease prevention approach by the

plant to defend itself against pathogens (Prell and Day, 2001). Inter-

development of fungus-resistant varieties (Rousseau et al., 2013).

specific hybridization of grapevines began in the 19th Century and

Fungus-resistant (FRG) result from interspecific cross-

was initially aimed at introducing pest and disease resistance in

breeding between the Mediterranean species V. vinifera and North

offspring (Galet 1999; Avenard et al., 2003; Reynolds, 2015). Many

American and Asian Vitis spp. such as V. riparia, V. amurensis and

FRG developed at that time carried undesirable “foxy” flavors in

V. rupestris that carries high resistance to fungal diseases, includ-

their wine, a default that was later attributed to the presence of

ing powdery and downy mildews, and grey rot. The firsts FRG

V. labrusca in the breeding process, resulting in the near elimina-

varieties, issued from traditional breeding, carried a significant

tion of this species in recent breeding (Hemstad and Luby 2000;

percentage of non-V. vinifera species in their genetic and were

Sun et al., 2011a). Later, several breeding programs implemented

therefore considered as “interspecific hybrids” (Sivcevˇ et al., 2010).

worldwide led to the development of varieties showing different

Recently, marker-assisted selection combined with multiple back-

characteristics such as cold-hardiness, short/long growing season,

crossing with V. vinifera varieties allowed the development of FRG

and pest resistance (detailed review by Reynolds, 2015).

Please cite this article in press as: Pedneault, K., Provost, C., Fungus resistant grape varieties as a suitable alternative for organic wine

production: Benefits, limits, and challenges. Sci. Hortic. (2016), http://dx.doi.org/10.1016/j.scienta.2016.03.016

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Table 1

Estimated growing area for significant FRG varieties grown in Europe, North America, and Asia.

Color Variety Estimated growing area (ha) Countries Ref.

a

red 200–1000 France, Italy, Switzerland 1

Baco Noir 200–1000 Canada, United States 2–5

Frontenac 200–1000 Canada, China, United States 2–4, 6

Maréchal Foch 200–1000 Canada, United States, France 2–5

Marquette 200–1000 Canada, United States 2–4

Regent >1000 Austria, Belgium, Bulgaria, Czech Republic, 1

Denmark, England, Germany, Hungary, Italy,

Netherlands, Sweden, Switzerland

white >1000 France 7

Bianca >1500 Austria, Belgium, Czech Republic, Denmark, 1,7

Hungary, Italy, Moldova, Netherlands,

Romania, Serbia, Switzerland

Cabernet Blanc 200–1000 Austria, Germany, Switzerland 1

Frontenac Gris 200–1000 Canada, United States 2–4

Kunleány >1500 Hungary 1,7

La Crescent 200–1000 Canada, United States 3, 4

Vidal >1000 Canada 2–5

Villard blanc 200–1000 France 7

Zala Gyöngye >2500 Hungary 7

a

(1) Rousseau et al., 2013; (2) Association des vignerons du Québec, 2013 (Personal communication); (3) Tuck and Gartner, 2013; (4) Mount Kobau Wine Services, 2014;

(5) Grape Growers of Ontario, 2015; (6) Li et al., 2015; (7) Eurostat, 1999–2009.

The recent sequencing of the Vitis sp. nuclear genome signifi- varieties grown in six different European countries, the number

cantly contributed to the implementation of molecular breeding of fungicide treatments was reduced by 73% and 82% in organic

strategies in grapevines (see the review by Di Gaspero and Foria, vineyards with low and medium disease pressure, respectively

2015). These tools assist breeders in the selection of parents or (Rousseau et al., 2013). In a survey involving 65 German vineyards

of the most promising offspring (Di Gaspero and Foria, 2015). So under organic management, growers reported having to spray FRG

far, many markers related to desirable agronomic and enological varieties 3.8 times per season on average (Becker, 2013).

characteristics (e.g., fruit ripening time, berry size, color, acidity, Estimations are that growing FRG varieties could cut production

aroma) have been identified in grapevine, including a number of costs by two in French vineyards (Galbrun, 2008). In California, it

markers for disease resistance (Emanuelli et al., 2011; Merdinoglu has been estimated that powdery mildew resistant varieties could

and Caranta, 2013; Di Gaspero and Foria, 2015; Zini et al., 2015). allow cost savings as high as 48 M$ per year for the production

Genes related to disease resistance contribute to the activation subsets of Crimson Seedless Table grapes, raisin grapes and Central

of pathogen-specific defense mechanisms (PSDM) that induce a Coast wine grapes (Fuller et al., 2014).

hypersensitive response in plant upon the detection of specific The disease resistance of FRG varies with cultivar’s genetic and

pathogen-encoded Avirulence proteins (Verhagen et al., 2010; Wu growing location (Pavlousekˇ et al., 2014). Therefore most FRG vari-

et al., 2014). PSDM are among the most effective defenses of plants eties show some susceptibility to different pathogens, including

against pathogens (Wu et al., 2014). The main genetic markers , powdery mildew, botrytis, black rot and anthrac-

related to PSDM in Vitis sp. include markers for downy mildew nose (Table 2). In organic management, these diseases are generally

(rpv1, rpv3, rpv10, rpv12) and for powdery mildew (ren1, ren3, run1) controlled using sulfur-based fungicides (i.e., Myco-San) (Rousseau

(Zini et al., 2015; Di Gaspero et al., 2012). et al., 2013; Siegfried and Temperli, 2008). When copper-based

So far, downy and powdery mildews, and botrytis have been pri- formulations are necessary, they are used at a much lower rate

marily targeted for the development of FRG varieties and several than for V. vinifera varieties (Van Der Meer and Lévite, 2010). In

cultivars bearing resistance to these diseases have been developed Québec (Canada), garlic powder suspension (marketed as Buran) is

over the last twenty years (Rousseau et al., 2013). Nevertheless, known among the local agronomists to efficiently prevent powdery

certain pathogens have been able to bypass the resistance mech- mildew in organic FRG.

anisms of certain FRG in field trials (Di Gaspero and Foria, 2015;

Merdinoglu and Caranta, 2013). Recently, the pyramidization of

resistance haplotypes from different grape species into new FRG

2.3. Yield

varieties prevented pathogens from bypassing resistance mecha-

nisms, therefore making FRG resistance more durable (Di Gaspero

The yield of organic V. vinifera grapevines can show 8–16%

and Foria, 2015; Merdinoglu and Caranta, 2013).

reduction compared to conventional grapevines (Bayramoglu and

Gundogmus, 2008; Guesmi et al., 2012). In contrast, FRG are

2.2. Economical and agricultural benefits of disease resistance often more vigorous and may show high productivity (10 000–20

000 T/ha), which could contribute to secure yields in organic pro-

The benefits of growing FRG cultivars go well upon the sector duction (Reynolds and Vanden Heuvel, 2009; Sun et al., 2011b;

of organic wine production. Indeed, most V. vinifera varieties have Rousseau et al., 2013; Barthe, 2015).

low to high susceptibility to fungal diseases that result in signif- Yields ranging from 7.0 to 11.8 T/ha were reported for FRG

icant production costs and economic losses (Fuller et al., 2014). grown under conventional management with minimal treatment

In Italy, the annual cost for controlling downy mildew in conven- in Switzerland (Siegfried and Temperli, 2008). In a five years study

tional vineyard typically ranges from 8 to 16 million Euros per carried out in State, organic grapevines

year, depending on disease pressure (Salinari et al., 2006). Under produced, on average, 30% less crop than vines grown using con-

medium disease pressure, 12 treatments per season are neces- ventional management (12.7 vs. 19.9 Tons/ha, respectively; Pool,

sary for traditional V. vinifera varieties grown under conventional 1995). Such reduction was caused by differences in soil quality (e.g.,

management (Rousseau et al., 2013). In a study including 183 FRG the organic plots was inferior to that of conventional plots) and by

Please cite this article in press as: Pedneault, K., Provost, C., Fungus resistant grape varieties as a suitable alternative for organic wine

production: Benefits, limits, and challenges. Sci. Hortic. (2016), http://dx.doi.org/10.1016/j.scienta.2016.03.016

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Table 2

Overview of disease susceptibility, vigor and hardiness/cold tolerance of 41 fungus-resistant varieties.

a,b ◦ c

Variety Susceptibility to diseases Vigor Hardiness/Cold tolerance ( F) Ref.

DM PM BOT BR AT DA CG EUT

reds

Baco noir + ++ + +++ + +++ ++ high −10 to −20 1–7

Baltica + + + moderate 20 to −30 1, 8–10

Cabernet Carbon /+ ++ −/+ 2, 5

Cabernet Jura − −/+ −/+ 2, 5

Chambourcin + + ++ +++ + ++ moderate/high −5 to −15 1–6, 11

Chancellor +++ ++ + + + +++ ++ + moderate −10 to −20 1–6, 12, 13

Chelois + +++ + + +++ ++ −10 to −20 6

Concord + ++ + +++ +++ + +++ high −15 to −25 3, 4, 6

De Chaunac ++ ++ + + + ++ ++ +++ moderate/high −15 to −25 1–5

Frontenac −/+ ++ ++ ++ + ++ high −20 to −30 1, 3, 4, 8–10

Léon Millot + ++ + + + ++ ++ high −15 to −25 1–7, 11

Lucie Kulhmann + ++ + + moderate 1, 12, 13

Maréchal Foch + ++ + ++ + +++ +++ moderate −15 to −25 1–7, 11, 14

Marquette − + + + +++ high −20 to −30 1, 8–10, 14

Petite Perle + + + + + moderate −20 to −30 1, 8, 9, 14

Regent + + + + −5 to −15 2, 5, 7, 15

Sabrevois + + + + moderate −20 to −30 1, 10, 13, 14

Skandia + + + ++ moderate −20 to −30 1, 8, 9, 14

St. Croix ++ ++ ++ + + moderate/high −20 to −30 1, 3, 6, 8–10, 12–14

whites

Aurore ++ ++ ++ +++ + ++ +++ high −10 to −20 3–7

Bianca + + −/+ ++ −5 to −15 2, 5, 7, 16

Bronner −/+ + + 5, 2, 11

Cabernet blanc ++ ++ /+ −5 to −15 2, 5

Frontenac blanc −/+ + ++ ++ high −20 to −30 1, 8–10, 14

Frontenac gris −/+ + ++ + high −20 to −30 1, 8–10, 14

Geisenheim + +++ + + moderate 1

Helios ++ + + 2, 5, 11

Hibernal ++ + + + + moderate 1, 5, 12, 13

Johanniter ++ −/+ ++ 2, 5, 11

Kunleány ++ +++ − 5, 16

La Crescent + + ++ ++ high −20 to −30 1, 3, 8–10, 14

Louise Swenson + + + + ++ low −20 to −30 1, 8–10, 14

Osceola Muscat ++ ++ + + + high −15 to −25 1, 8, 9

Seyval + +++ ++ ++ + + ++ + moderate −10 to −20 1–7, 10, 11, 13, 14

Solaris ++ −/+ ++ −10 to −20 2, 5, 11, 14

Soleil blanc + −/+ ++ 2, 5, 11

St. Pepin + + + + moderate −20 to −30 1, 6, 10, 12–14

Traminette + + + + +++ ++ ++ + moderate/high −10 to −20 1, 3, 4, 10

Vandal-Cliche + + + +++ moderate/high −15 to −25 1, 10, 12, 13

Vidal + ++ −/+ + + + +++ + moderate −5 to −15 1–6, 10, 12, 13

Villard blanc − + + 5, 15

a

DM: downy mildew; PM: powdery mildew; BOT: Botrytis; BR: black rot; AT: anthracnose; DA: dead arm; CG: crown gall; EUT: Eutypa.

b

−/+ = resistant and few susceptible according to several references; − = resistant; + =slightly susceptible; ++ = moderately susceptible; +++ = highly susceptible.

c

(1) Dubé and Turcotte, 2011; (2) Sivcevˇ et al., 2010; (3) Dami et al., 2005a; (5) Rousseau et al., 2013; (6) Reisch et al., 1993; (7) Lisek, 2010; (8) Provost et al., 2012b; (8)

Wolf, 2008; (9) Provost et al., 2013; (10) Carisse and Lefebvre, 2011; (11) Tuchschmid et al., 2006; (12) Khanizadeh et al., 2008; (13) Khanizadeh et al., 2005; (14) Plocher

and Parke, 2008; (15) Avenard et al., 2003; (15) Daniela et al., 2013; (16) Pavlousek,ˇ 2013.

the competition caused by cover crops in the organic plot (Pool, The quality of FRG wines has been a key topic since their devel-

1995). opment in the 19th Century. Back then, many French-American

hybrids were thought to produce wine of “satisfactory commer-

cial quality” and were winning medals in wine competitions (Paul,

1996a). Recent studies showed that the quality of FRG wines is

2.4. Marketing and wine quality

generally rated as equivalent to that of V. vinifera (Van Der Meer

and Lévite, 2010; Pedneault et al., 2012; Rousseau et al., 2013).

FRG varieties are nearly completely absent from wine market-

For example, in a blind tasting carried out on 52 FRG wines from

ing in major wine producing countries, a situation that limits their

Europe, 62% red FRG varieties (24 wines tasted), including Caber-

expansion because they remain unknown to consumers. In a survey

net Jura (VB 5-02), Cabertin (VB 91-26-17) and the old interspecific

of 255 wineries growing FRG varieties (25% of them being under

hybrid Chambourcin (J. Seyve 26-205), were noted as equivalent

organic management), 40% of respondents pointed the “problem

or superior to Merlot (reference wine), and 31% of white (28 wines

of unknown varieties” as the biggest handicap for the marketing

tasted) were classified as equivalent or superior to the reference

of FRG wines (Becker, 2013). As FRG varieties carry non-V. vinifera

Chardonnay wine, including the interspecific varieties Gf. GA.47-42

genes (even at low levels), they may suffer from the perception that

( Weiss X Seyval blanc), Saphira (Gm 7815-1), and

interspecific hybrids produce low-quality wines (Fuller et al., 2014).

(Fr 240-75) (Rousseau et al., 2013).

Similarly, organic wine has suffered from a reputation of aver-

A consumer study conducted in Switzerland concluded

age quality until recently (Collective, 2008). The need to educate

that 70–90% of consumers noted Solaris and Maréchal Foch

consumers to organic wines made from FRG varieties is therefore

wines as equivalent to V. vinifera and Zweigelt wines

significant.

Please cite this article in press as: Pedneault, K., Provost, C., Fungus resistant grape varieties as a suitable alternative for organic wine

production: Benefits, limits, and challenges. Sci. Hortic. (2016), http://dx.doi.org/10.1016/j.scienta.2016.03.016

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(used as reference wines), respectively, and 23–30% of consumers 3.2. Impacts of canopy management

judged the FRG wines as “clearly superior” to the reference V.

vinifera wines (Van Der Meer and Lévite, 2010). A consumer survey 3.2.1. Managing yield and berry quality

comparing 21 red FRG wines produced in Eastern Canada to three Controlling yield may contribute to enhance berry and wine

imported V. vinifera wines described as major selling products in quality, especially when the growing season is unfavorable to opti-

this area showed that 76% of FRG wines were judged as equivalent mal berry ripening (Berkey et al., 2011). Studies show that either

or superior to the reference wines (Pedneault et al., 2012). Most cluster or shoot thinning contribute to increased TSS and pH in FRG

FRG wines were blends that included or Frontenac, berries, but results vary from one year to another and, in many

with other locally grown FRG varieties (Pedneault et al., 2012). trials, harvest date had a higher impact than yield management

(Tables 9 and 10). For example, 36% yield reduction using cluster

thinning negatively impacted increased the level of C6 compounds

3. Growing FRG varieties for organic wine production

in juice of Seyval blanc berries during a hot growing season in

Quebec (Canada) (Pedneault et al., 2015). Reducing yield signifi-

The chemical composition of FRG is highly variable from one

cantly increased skin softness in Seyval blanc berries, suggesting

variety to another, which is attributable to their large genetic pool.

that it may impact physical barriers protecting berries from fungal

This entails the need to optimize the relationship between varieties

infections (Barthe, 2015).

and growing conditions. Knowledge of berry chemical composition

Geneva Double Curtain (GDC) training contributed to increase

and of the impact of canopy management practices is essential to

yield but conversely reduced TSS and pH in a season-dependent

achieve that goal. Studies cited in the following Sections (3.1 and

manner in (Reynolds et al., 1995). GDC and Umbrella-

3.2) were generally conducted under conventional management

Kniffin (UK) increased yield in Frontenac and Marquette,

and in northern areas; this is where most FRG have been grown and

respectively, but as UK increased TA and lowered TSS and pH in

studied. To our knowledge, no studies are available yet on canopy

Marquette, GDC increased TSS and pH, and lowered TA in Fron-

management practices for European PIWI varieties. Studies com-

tenac when compared to other training systems (Bavougian et al.,

paring organic to conventional management in V. vinifera varieties

2012; Martinson and Particka, 2013). Both Vertical Shoot Position-

have been reviewed by Provost and Pedneault (2016) and show that

ing (VSP) and High-Cordon (HC) increased the level of free volatile

organic management has generally a limited impact on berry and

terpenes in juice compared to GDC, Smart-Dyson (SD)

wine composition, and wine quality.

and Scott-Henry (SH) (Ji and Dami, 2008).

3.1. Berry and juice composition

3.2.2. Impact on wine

Shoot thinning significantly decreased yield and the level of C6

3.1.1. Juice

alcohols in Maréchal Foch wines, but did not impacted wine sen-

An overview of juice characteristics (e.g., TSS, pH, TA) of FRG

sory perception (Sun et al., 2011b). In contrast, taste and flavors

grown in different locations is presented in Table 3. Certain FRG

of wines derived from un-thinned control vines were preferred

varieties present high pH and high TA, which causes issues with

over those derived from cluster-thinned vines in Vandal-Cliche

microbial spoilage and color stability in wines (Morris et al., 1984a).

(Pedneault et al., 2015).

YAN level also varies largely among FRG varieties. The analysis

Chancellor wines issues from the GDC scored higher on berry

of 30 FRG varieties across Midwestern and Eastern United States

flavor than those issued from the Y-trellis system, whereas wines

showed YAN concentrations ranging between 89 and 938 mg/L

issues from Hudson River Umbrella (HRU) scored higher on color

(Stewart and Butzke, 2012). High YAN values (≥250 mg/L) are com-

and lower on earthy note intensities (Tables 9 and 10; Reynolds

mon in V. riparia-based FRG such as Frontenac (Mansfield, 2015a;

et al., 2004). GDC increased yield and decrease TSS and pH in Seyval

Slegers et al., 2015; Stewart, 2013). In addition, significant vari-

blanc, but GDC wines rated higher on melon notes intensity, and

ations are observed between years, growing areas and cultural

lower on earthy and vegetal flavors and astringency when com-

practices (Stewart, 2013).

pared to wines from 6-arms Kniffin (6AK) and Y-trellis systems

(Reynolds et al., 2004).

3.1.2. Berries

In Traminette, increasing cluster sunlight exposition improved

Red FRG berries typically show anthocyanin concentrations

wine color and sensory ratings for linalool, rose and spice aromas

ranging from 0.5 to 1.5 mg/g M3 G eq. FW in berries with low tan-

(Bordelon et al., 2008; Skinkis et al., 2010). Increasing cluster expo-

nin concentration ranging from 0.07 to 0.95 mg/g berry in seed, and

sition had little impact on aroma intensity in Seyval blanc wines,

from 0.03 and 0.79 mg/g berry in skin (Tables 4 and 5 ). Conversely,

but wines from exposed clusters were rated as superior (Reynolds

tannin levels in V. vinifera cv. Pinot noir reach 1.2 mg/g berry in

et al., 1986).

seed and 0.56 mg/g berry in skin (Springer and Sacks, 2014). The

profile of other phenolic compounds such as flavonols and phenolic

3.3. Challenges in FRG wine production

acid derivatives varies significantly from one FRG variety to another

(Tables 6 and 7). In red European PIWI varieties such as Caber-

Recent findings (Manns et al., 2013; Slegers et al., 2015; Springer

net Cortis and Regent, isorhamnetin-3-o-rutinoside is the major

and Sack, 2015) show that many aspects of current enology knowl-

flavonol whereas quercetin-3-o-glucoside is the major flavonol in

edge may have limited application with FRG varieties, because most

Baco noir and Lucy Khulmann (Ratnasooriya et al., 2011; Ehrhardt

of them present particular biochemical characteristics. Therefore,

et al., 2014). Caftaric acid is the major hydroxycinnamic derivative

particular winemaking practices should be developed for these

in most FRG varieties (Zhu et al., 2012; Manns et al., 2013; Ehrhardt grapes.

et al., 2014).

Stilbenes such as resveratrol are known to be involved in plant

3.3.1. Juice extraction and methanol

defense mechanism against fungal infections (Ehrhardt et al., 2014).

Many FRG varieties contain high levels of pectin that necessitate

trans-Resveratrol and its glucoside are the main stilbenes reported

the use of enzymes to increase juice yield at pressing. High pectin

in berries of French hybrid FRG varieties grown in Canada, whereas

levels have been thought to increase methanol concentration in

high levels of trans- and cis-piceid, pallitol and astringin have been

FRG wine (Lee et al., 1975). However, surveys showed that the level

reported in PIWI varieties from Italy and Germany (Table 8).

of methanol in FRG wines ranges between 20 and 197 mg/L, which

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6 K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx

Table 3

Overview of TSS, TA, pH and YAN levels in the juice of 41 fungus-resistant grape varieties from different growing areas.

Variety Localization TSS (Brix) TA pH YAN (mg/L) Ref.

Country Area Value Units Eq.

reds

a

Baco noir Canada Nova Scotia 18.5 0.82 % tartaric acid 2.66 1

Baltica Canada Québec 24.9 6.96 g/L tartaric acid 3.05 179 2

Cabernet Carbon Switzerland Stäfa 22.5–22.9 7.8–8.3 g/L n.a. 3.0–3.1 3

Switzerland Wädenswill 21.0 10.7 g/L n.a. 3.0 4

Cabernet Cortis Switzerland Wädenswill 22.3 10.6 g/L n.a. 3.1 4

Cabernet Jura Switzerland Wädenswill 21.2–24.0 8.1–10.2 g/L n.a. 3.0–3.4 3

Chambourcin China Beijing 20.2 4.2 g/L H2SO4 n.a. 5

Switzerland Wädenswill 19.0–21.6 12–12.4 g/L n.a. n.a. 4

United States 20.6–21.8 7.2–7.9 g/L n.a. 3.23–3.34 6

United States Ohio 20.1–21.3 10.7–11.2 g/L n.a. 3.20–3.27 7

Chancellor Canada British Columbia 18.5–23.5 12.2–16.0 g/L n.a. 3.12–3.37 8

United States Arkansas 14.8–17.1 0.74–0.93 % tartaric acid 3.47–3.79 9

Chelois United States Arkansas 14.8–18.2 0.68–1.07 % tartaric acid 3.61–3.80 9

Corot noir United States New York 15.0–17.8 7.5–11.6 g/L n.a. 3.34–3.75 213 10, 11

Frontenac Canada Québec 21.5–25.3 10.1–17.5 g/L tartaric acid 3.10–3.35 191–403 2,12–14

United States Iowa 21.4 10.4 g/L tartaric acid 3.39 15

United States Minnesota 22.9–26.4 14.9–18.5 g/L tartaric acid 2.86–3.12 16

United States Nebraska 19.7–22.5 14.6–20.9 g/L n.a. 2.92–3.12 17

Léon Millot Canada Nova Scotia 20.6 0.81 % tartaric acid 3.13 1

Switzerland Wädenswill 21.6 7.24 g/L n.a. 3.46 18

Lucy Khulman Canada Nova Scotia 21.1 0.76 % tartaric acid 3.11 1

Maréchal Foch Canada Nova Scotia 23.1 0.94 % tartaric acid 3.06 1

Canada Québec 21.6 7.24–10.3 g/L tartaric acid 3.18 95–108 12,14

Switzerland Wädenswill 21.6 9.7 g/L n.a. 3.2 4

United States Minnesota 24.6–26.1 8.7–11.8 g/L tartaric acid 2.99–3.05 16

United States New York 20.1–25.1 8.67–11.4 g/L n.a. 3.10–3.70 119 10, 19

Marquette Canada Québec 23.7–28.2 8.25–13.1 g/L tartaric acid 3.09–3.50 210–342 2, 12–14

United States Iowa 22.7 8.2 g/L tartaric acid 3.39 15

United States Minnesota 26.2–30.5 11.5–12.0 g/L tartaric acid 2.84–3.05 16

United States New York 21.9 9.37 g/L n.a. 3.09 329 10

Noiret United States New York 19.3 7.2 g/L tartaric acid 3.45 164 20

Petite Perle Canada Québec 22.9 7.38 g/L tartaric acid 3.33 299 2

Regent Germany n.d. 20.1–20.3 8.0–8.3 g/L n.a. – 21

Switzerland Stäfa 20.3–22.0 6.1–8.4 g/L n.a. 3.3–3.7 3

Sabrevois Canada Québec 18.6–19.2 13.4 g/L tartaric acid 3.16 213–221 12–14

Skandia Canada Québec 29 6.84 g/L tartaric acid 3.69 351 2

St. Croix Canada Québec 19.4–22.7 5.14–9.0 g/L tartaric acid 3.21–3.39 129–167 2, 12, 14

United States Iowa 16.8 6.3 g/L tartaric acid 3.73 15

United States Minnesota 20.9–24.1 3.87–4.96 g/L tartaric acid 3.08–3.50 16

Villard noir United States Arkansas 15.1–17.0 0.75–1.00 % tartaric acid 3.63–3.73 9

whites

Adalmiina Canada Québec 19.4 7.56 g/L tartaric acid 2.99 183 2

Aurore United States Arkansas 17.0–19.0 0.64–0.80 % tartaric acid 3.68–3.85 9

Bronner Switzerland Wädenswill 19.0–22.5 7.3–9.6 g/L n.a. 3.1 4

Cal 6-04 Switzerland Bevaix 21.0–24.5 10.2–12.4 g/L n.a. 2.9–3.0 3

Frontenac blanc Canada Québec 26.2 9.77 g/L tartaric acid 3.27 372 2

Frontenac gris Canada Québec 26.8 9.91 g/L tartaric acid 3.23 369 2

United States Minnesota 23.4–27.3 14.3–16.1 g/L tartaric acid 2.93–3.08 16

Helios Switzerland Wädenswill 21.6 8.5 g/L n.a. 3.2 4

Johanniter Switzerland Wädenswill 20.5 9 g/L n.a. 3.2 4

La Crescent Canada Québec 22.8 14.2 g/L tartaric acid 3 130 2

United States Minnesota 22.7–25.8 13.2–14.4 g/L tartaric acid 2.88–2.99 16

United States Iowa 21.8 8.9 g/L tartaric acid 3.36 15

Noah China Beijing 19.9 6.9 g/L H2SO4 5

Osceola Muscat Canada Québec 23.5 8.72 g/L tartaric acid 3.08 163 2

Seyval Canada Québec 17.7–22.4 8.77–12.3 g/L tartaric acid 2.86–3.16 94–180 22

United States Arkansas 16.9–17.6 0.60–0.89 % tartaric acid 3.69–3.91 9

Solaris Switzerland Stäfa 26.2–26.9 5.7–7.3 g/L n.a. 3.2–3.5 3

Switzerland Wädenswill 24.9–30.4 5.8–9.5 g/L n.a. 3.3 4

Soleil blanc Switzerland Wädenswill 22.9 8.73 g/L n.a. 3.43 18

St. Pepin United States Minnesota 22.0–23.8 8.8–9.8 g/L tartaric acid 2.94–3.25 16

Traminette United States New York 20.7 7.9 g/L tartaric acid 3.18 95 20

Vandal-Cliche Canada Québec 17.3–19.7 11.2–13.4 g/L tartaric acid 2.85–2.92 99–162 22

Verdelet United States Arkansas 16.2–20.1 0.35–0.84 % tartaric acid 3.63–4.32 9

Vidal blanc United States Michigan 18.8–20.2 0.65–1.02 % tartaric acid 3.28–3.30 23

Villard blanc China Beijing 20.1 6.8 g/L H2SO4 5

a

(1) Ratnasooriya et al., 2011; (2) Provost et al., 2012a; (3) Siegfried and Temperli, 2008; (4) Tuchschmid et al., 2006; (5) Zhu et al., 2012; (6) Miller et al., 1996; (7) Dami

et al., 2006; (8) Reynolds et al., 1995; (9) Morris et al., 1984b; (10) Manns et al., 2013; (11) Sun et al., 2012; (12) Slegers et al., 2015; (13) Pedneault et al., 2013b; (14) Pedneault

et al., 2013a); (15) Vos, 2014; (16) Haggerty, 2013; (17) Bavougian et al., 2012; (18) Striby, 2006; (19) Sun et al., 2011b; (20) Nisbet et al., 2014; (21) Eibach and Töpfer, 2003;

(22) Barthe et al., 2012; (23) Wolpert et al., 1983.

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HORTI-6344; No. of Pages 21 ARTICLE IN PRESS

K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx 7 e Ref. 1 2 3 4 3 3 3 4 4 5 6 1 7 2 7 7

d

– – – 6.1 44.9–62.3 4.71–6.66 22.4–23.8 Pt-3,5-O- diglucoside Method a a b c b b b c c b d a e a e e

DW DW DW DW

a a g g g g

FW FW FW

FW FW

a b a c a c c

mg/100 mg/100 mg/100 mg/100 mg/L mg/L mg/L Units mg/g mg/g mg/kg mg/kg mg/kg mg/L mg/L mg/L mg/L 0.04 0.22 0.17 Pl-3,5-O- diglucoside

1.95 0.61 1.3 50.6 148–163 50.5–56.9 128–153 Mv-3,5-O- diglucoside anthocyanin

84.6 224–1138 671–897 479–919 4.64–8.34 0.994–1.26 Total

.

21.7 59.7 60.1 25.6–29.5 6.12–7.01 15.3–17.8 Pn-3,5-O- diglucoside 2013

- al., p

-  et

128 91 313 Mv-3-(6 coumaroyl)-O- glucoside

Manns

3.11 9.84 4.7 15.7–19.8 1.98–2.39 3.47–3.60 Cy-3,5-O- diglucoside (7)

;

2007

- al., p

-  et

HPLC-DAD. – – – 34.0–49.0 2.88–7.39 12.2–15.0 Dp-3,5-O- diglucoside

e)

; 4.44 34 36.4 Pn-3-(6 coumaroyl)-O- glucoside varieties. Prajitna

520nm (6) eq. ␭

;

grape at

nd 0.72 0.32 Pl-3-O- glucoside 2012

- p al., -

UV–vis

 et

Zhu 31.5 36 69 Pt-3-(6 coumaroyl)-O- glucoside

fungus-resistant

(5)

; 190 79 96 220 110 234 203 7.3 12.2–17.0 45.1–89.6 28.9–31.1 Mv-3-O- glucoside of

malvidin-3,5-diglucoside

2014

in wines

al.,

Spectrophotometry -

p et -

and d)



4.84 57 41.5 4.1 0.77–1.05 1.62–3.39 0.87–1.39 Pn-3-O- glucoside quantified 7.15 60 49.9 coumaroyl)-O- glucoside Cy-3-(6 berry)

Pelargonidin. Ehrhardt

are

Pl:

(4)

whole ;

UPLC–MS/MS;

c)

2011 (skin,

- 150 30.1 100 200 130 87 71.3 12.0–16.7 16.1–49.0 9.38–16.6 Pt-3-O- glucoside p Malvidin;

- al., diglucosides



et

Mv:

berries eq.;

40.9 76 164 Dp-3-(6 coumaroyl)-O- glucoside

LC–MS/MS; in

b)

Peonidin;

40 2.93 21 55 42 30 24.4 1.00–1.54 1.14–2.21 0.78–0.84 Cy-3-O- glucoside Ratnasooriya assay;

Pn:

(3)

; eq. concentration -acetyl)-O-

 eq.

malvidin-3-glucoside 2011b

Petunidin; in

72.2 15.6 29.9 Mv-3-(6 glucoside

precipitation

al., Dp-3-O- glucoside* 190 66.6 130 200 250 408 221 14.5–18.0 12.6–56.3 6.72–14.9

Pt:

et

anthocyanin

Sun protein

total quantified

(2)

;

Cyanidin;

USA USA USA USA USA USA USA USA Canada Canada Canada Canada

are

and

-acetyl)-O-  Cy: Italy OH, NY, NY, Origin NY, NY, NS, Italy NS, NS, NS, Germany China NY, NY, NY,

malvidin-3-glucoside malvidin-3,5-diglucoside 2012

in in

-3-(6 al.,

profile

1.5 2.87 3.12 Pn glucoside et

Foch Foch Foch

cortis

Sun Adams-Harbertson

noir noir

4 Millot Noir Khulman

Delphinidin;

Quantified Quantified Monoglucosides (a) (1) c a e berry wine Léon wine Chambourcin Maréchal Marquette Variety Cabernet Lucy Maréchal Regent Maréchal berry Baco Corot Variety skin Corot skin b d Anthocyanin

Table *Dp:

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8 K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx

is slightly higher than V. vinifera wines (26–111 mg/L) but signif- 3.3.2. Biogenic amines

icantly lower than the recommended limits of OIV for both reds Exogenous nitrogen sources are routinely used to ferment V.

≤ ≤

( 400 mg/L) and whites ( 250 mg/L) (Lee et al., 1975; Organisation vinifera varieties. In FRG carrying high YAN levels (see Section

Internationale de la Vigne et du Vin, 2011). 3.1.1), this enrichment may cause an unwanted rise in fermentation

temperature and contribute to increase the level of fusel alcohols,

undesirable biogenic amines (e.g., histamine) and carcinogenic

Table 5

Flavan-3-ols profile and total tannin concentration in berries (seed, skin, whole berry) and wines of fungus-resistant grape varieties. Data on Vitis vinifera (Pinot noir var.)

are presented for comparison purposes (from Springer and Sacks, 2014).

Matrix Variety Origin (+)-catechin (−)-epicatechin epigallocatechin (−)-epicatechin 3-O-gallate

seed

red Baco noir NY, USA

ON, Canada 98 106

Corot noir NY, USA

NY, USA

DeChaunac NY, USA

ON, Canada 135 78

Frontenac QC, Canada

Léon Millot NY, USA

Maréchal Foch NY, USA

ON, Canada 46 42

QC, Canada

Marquette QC, Canada

Noiret NY, USA

Sabrevois QC, Canada

St. Croix QC, Canada

Vincent ON, Canada 155 284

*

Pinot noir NY, USA

white Seyval ON, Canada 21 23

skin

reds Baco noir NY, USA

Corot noir NY, USA

NY, USA

DeChaunac NY, USA

Frontenac QC, Canada

Léon Millot NY, USA

Maréchal Foch NY, USA

QC, Canada

Marquette QC Canada

Noiret NY, USA

Sabrevois QC Canada

St. Croix QC Canada

*

Pinot noir NY, USA

berry

red Baco Noir NS, Canada 1700 1100 15 140

Cabernet cortis Italy 94.5 101

Léon Millot NS, Canada 860 600 11 99

Lucy Khulman NS, Canada 1700 780 11 88

Maréchal Foch NS, Canada 2100 990 15 99

Regent Germany 176 148 57.4

Italy 167 115 36.7

white Johanniter Italy 84.9 42.1

Phoenix Germany 231 187 121.7

Italy 64.2 16.5

Solaris Italy 81 111 12.7

wine

red Baco noir NY, USA

Chambourcin China 1.23 1.23 3.28

GA, USA 4.1–9.3 2.6–14.8

Corot noir NY, USA

NY, USA 19.3–36.8 16.4–32.8

NY, USA

Frontenac QC Canada

Maréchal Foch NY, USA

NY, USA 36−337 11.5–62.1

QC, Canada

Marquette NY, USA 13.9–30.0 5.55–14.4

QC, Canada

Sabrevois QC, Canada

St. Croix QC, Canada

Noiret NY, USA

*

Pinot noir NY, USA

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Table 5 (Continued)

Matrix epigallocatechin procyanidin procyanidin procyanidin procyanidin procyanidin total Units Method Ref. gallate B1 B2 B3 B4 C1 tannins

seed

b c

red 0.454 mg/g berry CE a 1

a

33 91 41 127 10 mg/100 g seed CE or B2E b 2

0.33–0.53 mg/g berry CE a 3

0.917 mg/g berry CE a 1

0.264 mg/g berry CE a 1

4 22 2 12 tr mg/100 g seed CE or B2E b 2

0.200 mg/g epicatechin eq. c 4

0.594 mg/g berry CE a 1

0.763 mg/g berry CE a 1

14 43 23 61 4 mg/100 g seed CE or B2E b 2

0.377 mg/g epicatechin eq. c 4

0.259 mg/g epicatechin eq. c 4

0.208 mg/g berry CE a 1

0.069 mg/g epicatechin eq. c 4

0.170 mg/g epicatechin eq. c 4

60 106 27 45 tr mg/100 g seed CE or B2E b 2

1.19 mg/g berry CE a 1

white 3 9 2 0 mg/100 g seed CE or B2E b 2

skin

reds 0.178 mg/g berry CE a 1

0.19–0.31 mg/berry CE a 3

0.34 mg/g berry CE a 1

0.178 mg/g berry CE a 1

0.051 mg/g epicatechin eq. c 4

0.221 mg/g berry CE a 1

0.248 mg/g berry CE a 1

0.030 mg/g epicatechin eq. c 4

0.055 mg/g epicatechin eq. c 4

0.785 mg/g berry CE a 1

0.076 mg/g epicatechin eq. c 4

0.192 mg/g epicatechin eq. c 4

0.559 mg/g berry CE a 1

berry

red 8.7 mg/kg DW d 5

# #

43.8 106 28.2 mg/kg FW berries e 6

7.5 mg/kg DW d 5

7 mg/kg DW d 5

8 mg/kg DW d 5

# #

54.7 64.7 30.9 mg/kg FW berries e 6

# #

65.3 93.7 45.5 mg/kg FW berries e 6

# #

white 37.8 50.2 23 mg/kg FW berries e 6

# #

66.8 59.3 38.4 mg/kg FW berries e 6

# #

19.3 22.6 91.7 mg/kg FW berries e 6

# #

3.6 113 17.9 mg/kg FW berries e 6

wine

red 49 mg/L CE a 1

mg/L CE d 7

2.5–9.6 mg/L CE f 8

42.1–63.6 mg/L CE a 3

mg/L f 9

113 mg/L CE a 1

85.5 mg/L epicatechin eq. c 4

83 mg/L CE a 1

mg/L f 9

120 mg/L epicatechin eq. c 4

mg/L f 9

145 mg/L epicatechin eq. c 4

200 mg/L epicatechin eq. c 4

97.3 mg/L epicatechin eq. c 4

354 mg/L CE a 1

358 mg/L CE a 1

*

Vitis vinifera.

#

Procyanidins B2 and B4 quantified together.

a

Monomers are quantified as CE; dimers and trimers are quantified as B2E.

b

Analytical method: (a) Adams-Harbertson protein precipitation assay; (b) HPLC-UV–vis; (c) HPLC-Fluorescence; (d) LC–MS/MS; (e) UPLC–MS/MS; (f) HPLC-DAD.

c

(1) Springer and Sacks, 2014; (2) Fuleki and Ricardo da Silva, 1997; (3) Sun et al., 2012; (4) Pedneault et al., 2014; (5) Ratnasooriya et al., 2011; (6) Ehrhardt et al., 2014;

(7) Zhu et al., 2012; 8) Auw et al., 1996; (9) Manns et al., 2013.

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production: Benefits, limits, and challenges. Sci. Hortic. (2016), http://dx.doi.org/10.1016/j.scienta.2016.03.016

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10 K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx d c Ref. 1 1 1 1 1 1 1 2 3 4 4 4 Ref. 1 2 1 1 1 2 2 2 2 2 2 3 4 4 4 3 3

c

b

Method a a a a a a a b c c c c Method a b a a b b b b b a c c c a a

FW FW FW FW FW FW FW DW FW DW DW a DW FW b FW FW FW FW FW

a b

a a a mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/L mg/L mg/L mg/L mg/L Units Units mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/L mg/L mg/L mg/L

4.81 ethyl gallate

1.63 0.93–1.30 1.41–1.68 0.57–0.82 protocatechuic acid

syringetin-3-glucoside 5.69 3.1 0.2

7.79 2.9–17.0 4.64–10.7 6.19–12.6 4.45–8.00 gallic acid rutin 0.65–7.720.82–2.97 2.07–4.68 mg/L mg/L

acid

ND-0.70 ND-0.16 quercetin 5.63 0.54–1.29 2.37 3.31

derivatives dihydroxybenzoic acid ND 0.02 ND ND 0.03 ND ND 0.60–1.61 ND-2.09 ND-1.47 Benzoic

varieties.

0.61–3.68 2.37–3.84 3.09–6.64 GRP (GAE).

eq.

grape

acid

3.96 1.75 2.12 4.93 1.79 1.57 1.79 1.09 0.54–1.76 1.48–2.65 1.23–1.33 fertaric acid

gallic isorhamnetin-3-o-rutinoside 50.9 87.2 163.1 nd nd nd nd

. mg/L

2.97 4.78 1.81 2.59 2.28 1.54 2.44 ellagic acid in fungus-resistant

2013

al.,

white et

0.07–2.02 ND-0.71 ND-0.37 coumaric acid ethyl ester and quantified

Manns are red

varieties.

(4)

; 0.05–1.17 ND-0.61 0.01–1.17 caffeic acid ethyl ester quercetin-3-o-glucuronide 20.9 60 18.9 10.3 24.7 6.44 10.9 some

grape

of

1996

.

derivatives

al.,

wines et

2013

25.7 30 8.72 6.44 6.49 0.9 6.93 5.76 nd-9.3 2.09–13.5 1.18–10.6 4.79–7.24 coutaric acid

acid

al., and

Auw

et

(3)

benzoic fungus-resistant ;

1.36–14.2 3.54–4.82 0.43–2.25 coumaric acid berries

Manns

in

2012

quercetin-3-o-glucoside 20 34.1 16 8 45.6 8.1 20.2 17.2 16 32.7 6.84 1.49 ND

white (CAE);

(4)

al., ;

eq. and

et

ND-0.25 ND-0.37 ND cinnamic acid content

1996

red

acid

Zhu

al.,

(3)

et

; some

caffeic

54.6 36 12.7 17.9 26.3 1.02 15.5 0.7–15.3 7.67–30.4 26.7–58.8 caftaric acid of

derivatives Auw

2014

mg/L

(3) acid al.,

wines ; HPLC-DAD.

acids in

et

(c) kaempferol-3-o-glucoside 3.18 4.04 nd 5.38 4.42 nd 3.65

and

2012

al., HPLC-DAD. benzoic

derivatives

(GAE). c) et (QE).

Ehrhardt

quantified

USA USA USA berries

Canada Canada CanadaCanada 35

and and Hydroxycinnamic caffeic acid 2.2 4.40–11.50 2.98–6.64

eq. in eq.

(2)

are Zhu

Origin NS, NS, NS, Germany Italy Germany Italy China NY, NY, China China Italy Italy

;

acid (2)

; USA USA USAUSA 2.79–4.02 8.52–44.1

2011

Foch Foch

HPLC-ESI–MS/MS; cortis Italy

content

Italy Italy GA, Origin Italy Germany Italy Germany Italy China NY, NY,

al., quercetin gallic

blanc

2014

(b)

UPLC–MS/MS; noir NY,

Millot

Noir Khulman NS, derivatives et

b) al.,

Foch NY, mg/L mg/L cortis

et flavonol hydroxycinnamic

Variety Baco Cabernet Johanniter Chambourcin Noah Leon Lucy Marechal Regent Solaris Corot Maréchal Marquette Villard

in in

noir

the the

of of white Johanniter Phoenix Solaris Corot Maréchal Marquette Variety red Cabernet Regent red Chambourcin

red red white white

Ratnasooriya Ehrhardt

LC–MS/MS; UPLC–MS/MS;

6 7

Quantified (a) (1) Hydroxycinamic Quantified (a) (1) c c a a wine Matrix berry Matrix berry wine b b d Table Table

Overview Overview

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G Model

HORTI-6344; No. of Pages 21 ARTICLE IN PRESS

K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx 11 c Ref. 1 2 1 1 1 2 2 2 2 2 2 3 3 4 5 3 3 3 3 3 3 3 3 3 3 4 4 3 3 3

b

Method a b a a a b b b b b b c c a d c c c c c c c c c c a a c c c

a DW FW DW DW DW FW FW FW FW FW FW

RE RE RE RE RE RE RE RE RE RE RE RE RE RE RE RE RE RE

Units mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/kg mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L mg/L

astringin 0.26 4.01 0.28 0.13 0.15 0.18 0.13 0.16 0.18 0.30 0.10 0.17 0.22 0.10 0.16 0.04 0.35 0.13 0.30 0.04 0.12 0.06

pallidol 0.22 5.62 0.48 0.16 nd 1.94 0.26 0.26 0.14 0.36 0.09 0.40 0.28 0.38 0.16 0.06 0.315 0.25 0.25 0.11 0.24 0.19

purposes.

piceid

total 3.57 3.95 7.02 0.96 1.13 2.84 2.36 1.64 0.64 2.81 1.34 2.88 1.29 1.56 1.72 comparison

for

shown

are

-piceid , 0.41 11.3 0.44 0.18 0.57 0.16 nd nd nd nd cis (2007)

al.

et

-piceic .

Naugler

0.23 7.05 0.21 0.08 0.31 0.08 0.09 tr 9.3–29.6 0.16 0.5 trans 2007

from

al.,

et

wines,

Prajitna -resveratrol

vinifera (5)

; 9.8 9.0 8.5 7.8 trans glucoside Vitis

2012

on

al.,

et

Data

Zhu

-resveratrol (4)

; cis 0.01 0.17 0.01 0.01 0.02 nd nd 3.68 0.80 0.5–1.9 0.41 0.37 0.38 0.80 0.95 0.68 0.74 1.76 0.65 1.0 0.44 0.45 0.58 varieties.

2007

grape

al.,

et

-resveratrol

Naugler

trans 4.5 nd 4.3 4.3 4.7 1.94 nd nd nd nd nd 0.86 0.90 1.4–6.4 0.65 0.49 0.75 1.17 1.92 0.93 0.15 4.55 0.99 1.10 0.56 0.56 0.62 (3)

; fungus-resistant

of

2014

USA Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada Canada

al.,

wines

Origin NS, Italy NS, NS, NS, Germany Italy Germany Italy NS, China NS, NS, NS, NS, NS, China China NS, NS, NS, Italy Italy NS, OH, NS, NS, NS, NS, NS, et

HPLC-DAD. and

(c)

(RE).

Ehrhardt

berries

eq. Kuhlman

in

;(2)

reds 2011

Foch Foch

cortis

UPLC–MS/MS;

content

blanc

al.,

resveratrol

noir/Cabernet noir Kuhlman (b) Millot Millot Millot/Lucie

Noir noir

Khulman

et

Chaunac

vinifera

mg/L

Variety red Baco Cabernet red Baco Zweigelt/Cabernet Leon Lucy Maréchal Regent white Johanniter Phoenix Solaris Chambourcin de Leon Leon Lucie Maréchal white Noah Villard V. Pinot Pinot stilbene

in

the

of

berry

Ratnasooriya HPLC–MS/MS;

8

Quantified (a) (1) c a Matrix whole wine b Overview

Table

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Table 9

Impact of cultural practices on grapevine, berries, juices and wine from red varieties.

treatment variety # year main effects observed on ref.

grapevine berry/juice wine

a

cluster thinning Chambourcin 2–4 yield increased with crop thinning increased TSS, 1

level; TA, and pH

thinning favored

lignification and reduced

bud cold injury

Chambourcin 5 thinning reduced yield, thinning increased TSS, 2

increased cluster and berry and pH

weight

DeChaunac 3 same yield and berry no impact on TA, TSS, 3

weight and pH

DeChaunac 2 thinning increased TSS 4

and total phenols

Marquette nd thinning reduced yield thinning increased TSS 5

and pH

cluster thinning, Baco noir 3 pruning reduced yield, pruning increased pH 6

pruning cluster weight and TSS

cluster thinning, shoot Chancellor 3 thinning increase cluster thinning increased TSS, 7

thinning and berry weight pH, and red color

shoot thinning Concord nd thinning increase cluster thinning increased TSS 8

weight

Frontenac 1 similar cluster weight, 9

reduced berry weight

Frontenac 1 6 shoot/ft had highest no impact on TA, TSS, 10

cluster number and yield and pH

Maréchal Foch 2 thinning reduced yield, thinning increased TSS thinning decreased the 11

increased berry weight in juice, and berry level of C6 alcohols anthocyanins (cis-3-hexenol,

trans-2-hexenol,

1-hexanol) in wines

but impact on wine

sensory perception

were minor;

no impact on wine anthocyanin

b

training system Chancellor 5 GDC and YT increased When significant, GDC 12

yield, reduced cluster showed TSS, TA and pH

weight and showed in the lowest range,

optimal level of pruning and level of

weight when compared to anthocyanin in the

HRU, 6AK, MWC highest range

Chancellor 4 GDC and 6AK had highest 6AK and YT had the 6AK wines had lowest% 13

yields; 6AK and YT had lowest TSS and the ethanol, TA and pH;

highest cluster weight highest pH GDC wines had the

highest berry flavour

and YT the lowest;

HRU wines had the

highest color intensity

and the lowest

intensity in earthy

notes

no impact on wine

anthocyanins

Frontenac 2 GCD increased yield GDC had higher pH and 14

TSS, and lower TA

Frontenac nd VSP had higher total 15

phenol concentration

than SD

Marquette nd UK increased yield, cluster VSP increased TSS and 16

weight, and clusters/vine pH;

UK had lowest TSS and

pH, and higher TA

training system and Marquette 2 TWC had higher yield and TWC had lower TSS; 17

cluster thinning cluster weight cluster thinning

increased TSS in TWC

and VSP 18”

St. Croix 2 TWC increased yield and TWC had higher TSS; 17

cluster weight; cluster cluster thinning

thinning reduced yield increased TSS

a

(1) Dami et al., 2005b; (2) Dami et al., 2006; (3) Morris et al., 1987; (4) Wood and Looney, 1977; (5) Emling and Sabbatini, 2013; (6) Byrne and Howell, 1978; (7) Morris

et al., 2004; (8) Zabadal et al., 2002; (9) Rolfes et al., 2012; (10) Rolfes, 2014; (11) Sun et al., 2011b; (12) Reynolds et al., 1995; (13) Reynolds et al., 2004; (14) Bavougian et al.,

2012; (15) Bavougian et al., 2013; (16) Martinson and Particka, 2013; (17) Provost et al., 2012b.

b

6AK: 6-arm Kniffin; GDC: Geneva double curtain; HRU: Hudson River Umbrella; SD: Smart-Dyson; TWC: Top Wire Cordon; UK: Umbrella Kniffin; VSP: Vertical Shoot

Positioning; YT: Y-trellis.

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Table 10

Impact of cultural practices on grapevine, berries, juices and wine from white fungus-resistant grape varieties.

treatment variety # years main effects observed on ref.

grapevine berry/juice wine

a

cluster thinning Seyval 4 yield was similar no impact on TA, TSS, 1

between treatments; pH

thinning increased

berry weight

Seyval 1 thinning increased thinning increased TSS 2

cluster weight, reduced

yield

Vandal-Cliche 2 thinning reduced yield no impact on TA, TSS, 2

pH

Vidal 2 thinning increased thinning increased 3

cluster weight TSS/TA ratio

leaf removal Frontenac gris nd no impact on berry size no impact on TSS, TA, 4

pH

leaf removal, La Crescent nd no impact on berry size leaf removal + no 4

cluster thinning thinning increased TSS

cluster exposition vs Seyval 2 higher occurrence of higher TSS and lower non significant 5

shading bunch rot in shaded TA in exposed clusters differences between

clusters intensity of aroma,

perceived acidity and

quality of wines, but

authors stated that

wines from exposed

clusters were

“superiors”

Traminette 3 full shaded clusters had fruit exposure 6

3

lower TSS and PVT in increased wine color

juice; and sensory ratings for

fully exposed berries linalool, rose and spice

had higher PVT aromas

no impact on wine

taste, aftertaste and

mouthfeel

shoot and Seyval 2 shoot and cluster little impact on TSS, TA, wines issued from the 7

cluster thinning thinning reduced yield pH cluster thinning

treatment were

preferred by the

sensory panel only in

the cooler year;

under “normal”

conditions, panelists

rated the wines

similarly

shoot thinning La Crescent 1 similar cluster weight, 8

reduced berry weight

training system Kunleàny nd single curtain reduced 9

winter frosts

b

Seyval 4 GDC , 6-AK and YT had GDC and YT had lowest GDC wines had lowest% 10

highest yields and TSS and pH ethanol and TA;

lowest cluster weight GDC wines had higher

intensity on melon

notes, had low earthy

and vegetal character,

and had the lowest

astringency

Traminette 5 SH had highest yield SH and HC had higher 11

c

compared to HC TA, no impact on FVT

and PVT

Traminette 2 GDC had highest berry VSP, SH and SD had the 12

weight compared to lowest TA;

SH, SD and HC VSP had the highest

juice pH;

VSP and HC increased

the level of FVT in juice

training system Louise Swenson 2 cluster thinning TWC had higher TSS 13

and cluster thinning significantly reduced compared to VSP;

yield cluster thinning

increased TSS level in must

a

(1) Morris et al., 1987; (2) Barthe et al., 2012; (3) Wolpert et al., 1983; (4) Portz et al., 2010; (5) Reynolds et al., 1986 (6) Skinkis et al., 2010; (7) Berkey et al., 2011; (8)

Rolfes et al., 2012; (9) Balogh, 1989; (10) Reynolds et al., 2004; (11) Bordelon et al., 2008; (12) Ji and Dami, 2008; (13) Provost et al., 2012b.

b

6AK: 6-arms Kniffin; GDC: Geneva double curtain; HC: High cordon; SD: Smart-Dyson; SH: Scott Henry; TWC: Topwire Cordon; YT: Y-trellis.

c

FVT: free volatile terpenes; PVT: potentially-volatile terpenes.

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ethyl carbamate in wine (Vincenzini et al., 2009). Studies compar- and/or C6 compounds such as hexanol and cis-3-hexenol in wine

ing the level of biogenic amine in FRG and V. vinifera wines showed (Mansfield and Vickers, 2009; Pedneault et al., 2013a). The level

contradictory results (Baucom et al., 1986; Soleas et al., 1999), sug- of methoxypyrazines has been showed to decrease significantly in

gesting that large differences exist between varieties. Similarly, Frontenac berries during berry ripening (Pedneault et al., 2013a).

studies showed higher level of biogenic amines in organic com- The C9 aldehydes nonanal and trans,cis-2,6-nonadienal have been

pared to conventional wine (reviewed by Provost and Pedneault, shown to reach above-threshold concentrations in red FRG wines

2016). from Eastern Canada, suggesting that they could contribute to the

green notes found in certain FRG wines (Slegers et al., 2015). The

3.3.3. Tannins and color concentration of trans,cis-2,6-nonadienal has also been showed

In contrast with V. vinifera, FRG wines contain high levels of to increase in Frontenac and Marquette berries during ripening

diglycosylated and acetylated anthocyanins and high levels of (Pedneault et al., 2013a).

delphinidin-3-O-glucoside and petunidin-3-O-glucoside that may

provide their wines with purple-blue colors (Table 4) (Manns and

3.4. Improving FRG wine quality

Mansfield, 2012; Manns et al., 2013). This trait varies largely among

FRG varieties.

Historically, assays carried out to improve the quality of FRG

Color instability and lack of mouthfeel are major issues in red

wine were primarily meant to reduce the occurrence of foxy fla-

FRG wines (Manns et al., 2013; Springer and Sacks, 2015). FRG

vors with the use of different winemaking processes. In 1974,

wines generally carry low tannin levels (≤200 mg/L CE eq.) and

carbonic maceration was found to efficiently reduce foxy flavors

high anthocyanin levels (200–1200 mg/L M3G eq.) (Tables 4 and 5).

in red Concord (V. labrusca) wines (Fuleki, 1974; Table 11). Car-

In comparison, red V. vinifera wines contain between 150 and

bonic maceration (CM) was initially developed to reduce oxidation

600 mg/L CE eq. of tannins and between 200 and 400 mg/L of antho-

reactions occurring spontaneously in grapes in order to preserve

cyanin, depending on the variety and winemaking process (Casassa

fruit flavors (Paul, 1996b). In organic wine production, restrict-

and Harbertson, 2014; Kilmister et al., 2014; Pedneault et al., 2014;

ing contact between berries and air using CM may be of particular

Springer and Sacks, 2014). Indeed, studies have demonstrated that

interest because organic grapes have been shown to carry twice as

the majority of traditional extraction processes for FRG winemak-

much polyphenol oxidase activity compared to conventional ones

ing lead to high amounts of anthocyanin rather than tannins, and

(Núnez-Delicado˜ et al., 2006).

yield very dark wines with little mouthfeel (Table 11, Auw et al.,

White FRG wines such as Chardonnel, Solaris and La Crescent

1996; Manns et al., 2013; Pickering and Pour Nikfardjam, 2007;

generally present desirable flowery notes that may be related

Pour Nikfardjam and Pickering, 2008).

to compounds such as C13-norisoprenoids (e.g., ␤-damascenone)

Poor tannin extractability as well as poor retention of exoge-

and monoterpenes (e.g., linalool) located in berry skin (Table 12;

nous tannins in FRG wine was recently related to high protein

Cadwallader et al., 2009; Savits, 2014; Liu et al., 2015). In fact,

concentration that may contribute to precipitate tannins dur-

extended skin maceration (24 h cold-soak and 30 h on-skin fermen-

ing winemaking (Mansfield, 2015b; Springer and Sacks, 2014).

tation) significantly improved the intensity of floral notes in Solaris

Recently, pathogenesis-related proteins have been found to inter-

wines, but also increased green vegetable notes (Zhang et al., 2015).

fere with tannin retention in FRG wines (Springer et al., 2016).

Conversely, short cold-soaks (3–8 h) did not improve the aroma

PR-proteins expressed in reaction to biotic stresses are the major

intensity of Traminette wine (Skinkis et al., 2010).

proteins in V. vinifera wines; they are resistant to proteases and

A recent study reported for the first time the presence of 3-

to low pH values, making them difficult to remove or denature

mercaptohexanol in the FRG variety Cayuga, at a concentration of

(Ferreira et al., 2004).

195 ng/L (Musumeci et al., 2015). This compound is a highly odor

The level of PR-proteins in wine is partly related to the disease

potent thiol (odor perception threshold: 60 ng/L) that produces a

pressure in the vineyard. For instance, organic V. vinifera wines

grapefruit aroma in (Musumeci et al., 2015). Cayuga

were found to contain higher levels of PR-proteins compared to

white is an offspring of Seyval blanc (Seyve-Villard 5–276), a variety

non-organic ones, which was attributed to their higher exposure

that has often been used during the breeding of recent FRG varieties.

to fungi in the field (Sauvage, 2011). The impact of disease pres-

This suggests that 3-mercaptohexanol could be present in other

sure and its possible modulation of PR-protein concentration in FRG

FRG varieties, although its presence has not yet been investigated.

varieties grown under organic management have not been studied

Based on this finding, viticulture (e.g., nitrogen status, disease con-

yet.

trol) and winemaking practices (e.g., oxidation control) that either

The solution currently proposed to resolve the tannin retention

enhance thiol production in berries, protect thiol during winemak-

issue in FRG wine is to treat the juice or the wine with bentonite in

ing, or lead to thiol expression in wine could contribute to increase

order to precipitate the proteins and then add exogenous tannins

the occurrence of tropical aroma in FRG wines (Musumeci et al.,

(Springer et al., 2016).

2015).

Blending is one of the most efficient ways to optimize the aroma

3.3.4. Aroma

of FRG wines. Indeed, most FRG varieties present a very large and

The majority of studies on the aroma of FRG wines focused on

rich flavor range, and many of them have complementary fla-

the well-known “foxy compounds” that provide the “hybrid char-

vor profiles (Slegers et al., 2015). Blending may also significantly

acter” to V. labrusca-based FRG wines. Recent GC-Olfactometry/MS

improve wine balance, especially acidity, reduce bitterness and

analyses demonstrated that most foxy compounds such as o-

improve wine’s overall bouquet. Such richness makes FRG varieties

aminophenone and methyl anthranilate are not very abundant in V.

suitable to a large range of styles that have a high potential to appeal

riparia based FRG wines (Sun et al., 2011a). Similarly, no foxy com-

to consumers.

pounds were reported from GC-O/MS analyses of Frontenac wines

from Minnesota (Mansfield and Vickers, 2009; Table 12).

Many red FRG wines are known for their fruitiness but 4. Conclusion

herbaceous notes are also reported in certain varieties such

as Cabernet cortis, Prior, Regent and Frontenac, among others The well-documented susceptibility of V. vinifera cultivars to

(Table 12; Mansfield and Vickers, 2009; Rousseau et al., 2013). major diseases such as powdery mildew, downy mildew and botry-

Herbaceousness could relate to the presence of methoxypyrazine tis is a significant challenge in organic viticulture. Increasing the

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Table 11

Overview of the impact of several winemaking processes on the chemistry and the sensory perception of wines made from fungus-resistant grape varieties.

Treatments Variety Yeast strain Impacts on wine ref.

Basic chemistry Phenolic and volatile Sensory perception

compounds

Extraction and alcoholic fermentation processes

a

3 vs 8 h cold-soak at Traminette Lalvin K1-V1116 no perceptible impact 1

12 C on wine aroma

intensity

− direct press after Solaris Lalvin DV10TM 24 h cold soak with or extended skin contact 2

crushing without skin (24 h cold soak + 30 h

− whole cluster press fermentation increased skin fermentation)

6 h cold soak total polyphenols; increased the intensity

− 24 h cold soak extended skin contact of floral (Rose,

− 6 or 24 h cold (24 h cold soak + 30 h Elderflower) and green

soak + 30 h skin skin fermentation) vegetable descriptors.

fermentation increased the

*chemical concentration of

deacidification used on linalool, hotrienol,

all treatments ␣-terpineol,

␤-damascenone and

S-methyl thioacetate in

wine.

- cold-soak − enzyme Maréchal Foch GRE hot press increased the 3

addition at crush level of non

- tannins addition at anthonyanin phenolic

crush (including caftaric acid,

- hot press compared to coutaric acid, catechin,

skin fermentation epicatechin, catechin,

(7 days) rutin), and the level of

monoglucoside and diglucoside

anthocyanins;

tannin addition and

hot press increased

tannin concentration

but had no impact on

their mean degree of

polymerization.

Corot noir GRE tannin addition 3

increased tannin

concentration whereas

hot press decreased it,

both did not impact the

mean degree of

polymerization.

- hot press Marquette GRE hot press increased 3

- skin fermentation anthocyanin

(7 days) monoglucosides and

tannin concentrations.

- immediate press for Chambourcin Prise de mousse hot press (juice and 4

juice wine) increased color

- immediate press for intensity (A520 nm);

wine 13d and 21d skin

- hot press for juice fermentation decrease

- hot press for wine color intensity

- skin fermentation compared to 7d

(7, 13, and 21 days) treatment;

immediat press (juice

& wine) increased

browning (Hue);

extended on-skin

fermentation

decreased the level of

caftaric acid, coutaric

acid, and procyanidin

B3, and increased the

level of gallic acid;

hot press increased

total phenol

concentration.

-carbonic maceration Concord no starter in CM; carbonic maceration carbonic maceration 5

(CM) at 15 or 27 C, for 1 unknown starter used reduced the occurrence decreased the intensity

or 2 weeks in other trials of bluish tones, the of V. labrusca typical

- skin fermentation concentration in total flavor.

- hot press phenols and decreased

the concentration in

methyl anthranilate.

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Table 11 (Continued)

Treatments Variety Yeast strain Impacts on wine ref.

Basic chemistry Phenolic and volatile Sensory perception

compounds

two experiments: Maréchal Foch Epernay II the use of 100% whole 100% whole berries 6

1) Standard berries or whole and whole clusters

fermentation with clusters decreased the decreased total

varying percentage of level of residual sugars; phenols and color;

whole berries; 8d CM decreased TA extended CM (8d)

2) CM with whole and alcohol percentage, increased total

clusters (2, 4, 8 days) and increased pH phenols.

® ® ®

addition of OptiRed Baco noir EC1118 OptiRed increased the OptiRed had no 7,8

(inactivated yeast concentration of perceptible impact on

derivative) at the dimers procyanidins; wine mouthfeel.

®

beginning of skin OptiRed increased the

fermentation (19 days) concentration of

tyrosol and quercetin,

and decreased the

concentration of caffeic

acid, grape reaction

product, and delphinidin.

® ®

Maréchal Foch EC1118 OptiRed increased the OptiRed had no

concentration of dimer perceptible impact on

and trimers wine mouthfeel. procyanidins; ®

OptiRed increased the

concentration of caffeic

acid and decreased the

concentration of

caftaric acid,

p-coumaric acid, and

quercetin.

malolactic fermentation Chancellor, DeChaunac, Montrachet Non-inoculated control wines fermented with 9

(MLF): Maréchal Foch did not completed PSU-1 and LS-5A were

non inoculated control MLF; preferred over ML-34

compared to inoculation PSU-1 and LS-5A wines;

with the Leuconostoc strains completed MLF all inoculated wines

oenos strains ML-34, faster than ML-34; were preferred to the

PSU-1, and LS-5A MLF increased wine pH non inoculated control.

in Chancellor (strain

LS-5A only) and

Maréchal Foch,

decreased TA, and

increased volatile

acidity in all wines.

Post-fermentation processes and aging

French (Nevers; Seyval n.d. american oak increased perceptible changes in 10

Limousin) and American gallic acid content of sensory attributes of

oak barrels aging wine at a slightly wines compared to

compared to unoaked higher rate than French unoaked control were

control oak; perceptible after 7

the concentration of weeks of aging;

other non-flavonoid by the 12th week,

compounds wines were

(photocatechuic, distinguishable

vanillic, caffeic, according to the oak

syringic and type used for aging.

p-coumaric acids) were

no affected.

a

(1) Skinkis et al., 2010; (2) Zhang et al., 2015; (3) Manns et al., 2013; (4) Auw et al., 1996; (5) Fuleki, 1974; (6) Miller and Howell, 1989; (7) Pour Nikfardjam and Pickering,

2008; (8) Pickering and Pour Nikfardjam, 2007; (9) Giannakopoulos et al., 1984; (10) Jindra and Gallander, 1987.

use of FRG varieties would allow significant benefits for organic winemaking practices can contribute to improved FRG wine qual-

and conventional growers, including reducing the number of treat- ity. Further research on these topics are needed to provide a

ments per season, increasing grape yield and reducing labor costs. more comprehensive understanding of the optimal cultural and

In addition, FRG would allow significant reduction in the use of winemaking practices for FRG varieties, with an emphasis on the

copper-based fungicide, therefore contributing to decrease cop- potential side-effects from disease resistance that could interfere

per accumulation in vineyard soils, especially in areas under high with the development of high quality wines (e.g., PR-proteins).

disease pressure. The commercialization of organic FRG wines faces a double

Consumer surveys have showed that wines made from FRG challenge of growing grape varieties that are generally unknown

varieties are at least equivalent and frequently rated as supe- to consumers and doing it under organic management. Both FRG

rior to V. vinifera wines in terms of quality. Trials conducted over and organic wines previously suffered from the production of bad

the last 30 years have showed that canopy management and wines that contributed to negative opinions among consumers

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K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx 17

Table 12

Overview of volatile compounds, impact odorants and sensory descriptors in wines from 20 fungus-resistant grape varieties.

*

Variety Method Analytical Sensory Ref.

1

Volatile compounds Concentration (␮g/L) Aroma and flavors descriptors

reds

a

Cabernet Cortis sensory analysis using foxy, herbaceous, bell 1

similar descriptors for pepper, jammy, spicy

all varieties evaluated

Chancellor descriptive analysis berry, currant, earthy, 2 vegetal

**

Frontenac descriptive analysis & ethyl isobutyrate 0.1–0.5 mg/L cherry, black currant, 3

G C-O/MS vegetal, earthy

ethyl lactate 0.2–1.5 mg/L

ethyl 2-methylbutanoate 30−200

ethyl 3-methylbutanoate 40−720

1-hexanol 0-0.8 mg/L

ethyl hexanoate 1.2–3.6

phenethyl alcohol 0.8–2.1 mg/L

diethyl succinate 1.4–4.4

octanoic acid 0.8–5.4

ethyl decanoate 1.1–3.7

Frontenac GC–MS & odor activity nonanal 40 4 values

trans,cis-2,6-nonadienal 1.26

␤-damascenone 3.99

ethyl 2-methylpropanoate 281

ethyl hexanoate 450

ethyl octanoate 1128

Maréchal Foch GC–MS & odor activity nonanal 34.8 4 values

trans,cis-2,6-nonadienal 1.44

␤-damascenone 2.25

ethyl 2-methylpropanoate 281

ethyl hexanoate 227

ethyl octanoate 781

Marquette GC–MS & odor activity nonanal 52.2 4 values

trans,cis-2,6-nonadienal 1.15

␤-damascenone 2.47

linalool 36.2

ethyl 2-methylpropanoate 254

ethyl butanoate 327

ethyl hexanoate 803

ethyl octanoate 2383

Othello GC–MS & odor activity methyl anthranilate 5 values trans-2-hexenal hexanol

ethyl 3-methylbutanoate nonanal 2-phenylethanol

ethyl decanoate

Prior Sensory analysis using foxy, fusel alcohol, 1

similar descriptors for herbaceous, fresh fruit,

all varieties evaluated jammy

Regent Sensory analysis using foxy, animal, herbaceous, 1

similar descriptors for spicy

all varieties evaluated

Sabrevois GC–MS & odor activity nonanal 33.4 4 values

trans,cis-2,6-nonadienal 1.13

eugenol 23.1

ethyl 2-methylpropanoate 283

ethyl octanoate 643

St. Croix GC–MS & odor activity nonanal 30 4 values

trans,cis-2,6-nonadienal 1.19

eugenol 3.38

ethyl 2-methylpropanoate 265

ethyl octanoate 768

whites

Cayuga white GC-O/MS (Charm) 2-phenylethanol 6

␤-damascenone

C3 and C4 fatty acids

ethyl butyrate

linalool

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18 K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx

Table 12 (Continued)

*

Variety Method Analytical Sensory Ref.

1

Volatile compounds Concentration (␮g/L) Aroma and flavors

descriptors

GC–MS 3-mercaptohexanol 195 ng/L 7

Cabernet blanc Sensory analysis using herbaceous, citrus fruit, 1

similar descriptors for tropical fruit

all varieties evaluated

Chardonnel descriptive analysis, ethyl butanoate 147−422 fruit, floral, spicy 8

GC-O. GC–MS, and

OAVs

ethyl 3-methylbutanoate ND-25.1

ethyl hexanoate 721−999

3-methyl-1-butanol 37.5–170 ppm

2-phenylethanol 8.94–23.4 ppm ␤

-damascenone 1.4–3.2

sotolon detected in GC-O; not

quantified

Johanniter sensory analysis using sulfur, white fruit in syrup, 1

similar descriptors for dry fruit, apricot

all varieties evaluated

La Crescent descriptive analysis apricot, grapefruit, lychee, 9

pineapple, rose

Muscaris sensory analysis using sulfur, fusel alcohol, citrus 1

similar descriptors for fruit, tropical fruit, white

all varieties evaluated fruit in syrup

Seyval descriptive analysis floral, apple, earthy, melon, 2

vegetal

descriptive analysis fruity, vegetal, 10

caramelized, pungent

GC-O/MS (Charm) ␤-damascenone 6 2-phenylethanol

methyl anthranilate

ethyl 2-methylbutanoate

vanillin

Solaris descriptive analysis, ethyl butyrate peach/apricot, Muscat, 11

GC–MS melon, banana, strawberry

propyl acetate

butyl acetate

3-methylbutyl acetate

hexyl acetate trans-3-hexenyl-acetate

2-phenylethyl acetate

descriptive analysis fruity, citrus, tropical fruit, 12

flowery, elderflower

***

Vidal GC-O/MS & OAVs ethyl 2-methylbyturate 14.2 13

␤-damascenone 11.3

decanal 13.7

geranyl acetone 0.26

ethyl hexanoate 878

1-octanol 9.34

nerol oxide 6.98

4-vinylguaiacol 111

descriptive analysis young wines: apple, 14

tropical fruit, citrus

aged wines: cooked

vegetables, straw, oxidized,

pungent

GC-O/MS (Charm) ␤-damascenone 6 2-phenylethanol linalool

*

3 to 9 wines per varieties.

**

Compounds with frequency of detection higher than 75%. ***

Nose-perceptible odorants.

a

(1) Rousseau et al., 2013; (2) Reynolds et al., 2004; (3) Mansfield and Vickers, 2009; (4) Slegers et al., 2015; (5) Radulovic´ et al., 2010; (6) Chisholm et al., 1994; (7)

Musumeci et al., 2015 (8) Cadwallader et al., 2009; (9) Savits, 2014; (10) Andrews et al., 1990; (11) Liu et al., 2015; (12) Linden, 2014; (13) Bowen and Reynolds, 2012; (14)

Chisholm et al., 1995.

concerning these products. Therefore, significant efforts are neces- regarding the future of these varieties, in both organic and conven-

sary to demonstrate and improve the quality of organic FRG wines, tional viticulture.

and have them accepted by consumers. According to the Web of Sci-

ence database, the number of scientific publications involving FRG

Acknowledgements

varieties increased by 86% between 2011 and 2016, when compared

to the 2005–2010 period (search restricted to “fungus resistant

The authors wish to thank Gaëlle Dubé, viticulture specialist,

grape or varieties” and “interspecific or varieties”).

for valuable information on organic grape production from FRG

Such rise in the interest for FRG opens the door to great expectations

varieties, and Dr. Ross Stevenson (UQAM), for kindly reviewing the

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production: Benefits, limits, and challenges. Sci. Hortic. (2016), http://dx.doi.org/10.1016/j.scienta.2016.03.016

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K. Pedneault, C. Provost / Scientia Horticulturae xxx (2016) xxx–xxx 19

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